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Foundations-Of-Parasitology Gerald D. schmidt & Larry S. Roberts’ Foundations of parasitology Fascioloides magna, a very large liver fluke that parasitizes deer and cattle (p. 272). The body of this specimen was rendered transpar- ent, leaving only pigment in its intestinal tract visible. An enlarger was used to project an image directly onto photographic paper, which after development clearly shows the trematode’s highly branched gut. Courtesy of William C. Campbell eighth edition Gerald D. schmidt & Larry S. Roberts’ Foundations of parasitology larry s. roberts florida international university john janovy, jr. University of nebraska–lincoln GERALD D. SCHMIDT & LARRY S. ROBERTS’ FOUNDATIONS OF PARASITOLOGY EIGHTH EDITION Published by McGraw-Hill, a business unit of The McGraw-Hill Companies, Inc., 1221 Avenue of the Americas, New York, NY 10020. Copyright © 2009 by The McGraw-Hill Companies, Inc. All rights reserved. Previous editions © 2005, 2000, and 1996. No part of this publication may be repro- duced or distributed in any form or by any means, or stored in a database or retrieval system, without the prior written consent of The McGraw-Hill Companies, Inc., including, but not limited to, in any network or other electronic storage or transmission, or broadcast for distance learning. Some ancillaries, including electronic and print components, may not be available to customers outside the United States. This book is printed on recycled, acid-free paper containing 10% postconsumer waste. 1 2 3 4 5 6 7 8 9 0 QPD/QPD 0 9 8 ISBN 978–0–07–302827–9 MHID 0–07–302827–4 Publisher: Janice Roerig-Blong Executive Editor: Patrick E. Reidy Director of Development: Kristine Tibbetts Senior Project Manager: Kay J. Brimeyer Senior Production Supervisor: Kara Kudronowicz Outside Developmental Services: Margaret B. Horn Associate Design Coordinator: Brenda A. Rolwes Cover Designer: Studio Montage, St. Louis, Missouri (USE) Cover Image: Biomphalaria glabrata courtesy of Si-Ming Zhang, The University of New Mexico Lead Photo Research Coordinator: Carrie K. Burger Compositor: Lachina Publishing Services Typeface: 9.5/11 Times Roman Printer: Quebecor World Dubuque, IA Library of Congress Cataloging-in-Publication Data Roberts, Larry S., 1935- Gerald D. Schmidt & Larry S. Roberts’ foundations of parasitology / Larry S. Roberts, John Janovy, Jr. — 8th ed. p. cm. Includes index. ISBN 978-0-07-302827-9 — ISBN 0-07-302827-4 (hard copy : alk. paper) 1. Parasitology. I. Janovy, John, 1937- II. Title. III. Title: Gerald D. Schmidt and Larry S. Roberts’ foundations of parasitology. IV. Title: Foundations of parasitology. QL757.R585 2009 591.7'857—dc22 2008031997 www.mhhe.com about the authors ative accomplishments consist of approximately 90 scientific LARRY S. ROBERTS papers and book chapters; 14 books, including Keith County Journal, On Becoming a Biologist, Teaching in Eden, Out- Larry S. Roberts, professor witting College Professors, and Foundations of Parasitology emeritus of biology at Texas (with Larry Roberts); the screenplay for the televised version Tech University and an adjunct of Keith County Journal (Nebraska Public Television); and professor at Florida Interna- numerous popular articles. His teaching experiences include tional University, has exten- almost continuous service in the large-enrollment freshman sive experience teaching biology course; Field Parasitology (BIOS 487/887) at the parasitology, invertebrate zool- Cedar Point Biological Station; Invertebrate Zoology (BIOS ogy, marine biology, and de- 381); Parasitology (BIOS 385); a decade in BIOS 103/204 velopmental biology. He (Organismic Biology/Biodiversity); and numerous honors received his ScD in parasitol- seminars. He has supervised 18 MS students, 14 PhD stu- ogy at Johns Hopkins Univer- dents, and approximately 50 undergraduate researchers, in- sity and has co-authored cluding 10 Howard Hughes scholars. His honors include the Foundations of Parasitology University of Nebraska Distinguished Teaching Award from the first edition through (1970), University Honors Program Master Lecturer (1986), this, the eighth edition. He is American Health magazine book award (1987, for Fields of also co-author of Integrated Principles of Zoology, Biology of Friendly Strife), University of Nebraska Outstanding Re- Animals, and Animal Diversity, and is author of The Underwa- search and Creativity Award (1998), The Nature Conser- ter World of Sport Diving. vancy Hero recognition (2000), and the American Society of Dr. Roberts has published many research articles and re- Parasitologists Clark P. Read Mentorship Award (2003). views. He has served as president of the American Society of Parasitologists, Southwestern Association of Parasitologists, and Southeastern Society of Parasitologists, and is a member of GERALD D. SCHMIDT numerous other professional societies. He received the Henry Baldwin Ward Medal from the American Society of Parasitolo- Gerald D. Schmidt was pro- gists. Dr. Roberts also serves on the editorial board of the jour- fessor of biology at the Uni- Parasitology Research nal . His hobbies include scuba diving, versity of Northern Colorado underwater photography, and tropical horticulture. (UNC) when he passed away. Dr. Roberts can be contacted at: Lroberts1@compuserve He received his PhD from .com Colorado State University. He was active in research and promoting research activities JOHN JANOVY, JR. at UNC, and he published more than 160 research arti- John Janovy, Jr. (PhD Uni- cles in scientific journals, as versity of Oklahoma, 1965) is well as six books. He re- the Paula and D. B. Varner ceived awards from UNC for Distinguished Professor of outstanding teaching and for Biological Sciences at the distinguished scholarship. He University of Nebraska–Lin- was a board member of the World Federation of Parasitolo- coln. His research interest is gists; a Fellow of the Royal Society of Tropical Medicine parasitology, with particular and Hygiene, London; and a Fellow of the Royal Society of focus on parasite ecology and South Australia. life cycles. He has been direc- Dr. Schmidt served the American Society of Parasitolo- tor of the Cedar Point Biolog- gists as secretary-treasurer for seven years. He was co-author ical Station, interim director of Foundations of Parasitology through the first four edi- of the University of Nebraska tions. His hobbies were hunting and fishing, especially fish- State Museum, and an assis- ing, and he wrote a book on fishing. tant dean of Arts and Sci- Dr. Schmidt died on 16 October 1990; many more de- ences, and he is currently the secretary-treasurer of the tails of his life can be found in the Journal of Parasitology, American Society of Parasitologists. His scholarly and cre- 78:757–773. v brief contents 23 Nematodes: Trichinellida and Dioctophymatida, Preface xv Enoplean Parasites 399 24 Nematodes: Tylenchina, Pioneering 1 Introduction to Parasitology 1 Parasites 413 2 Basic Principles and Concepts I: Parasite 25 Nematodes: Strongyloidea, Bursate Systematics, Ecology,and Evolution 11 Rhabditidans 419 3 Basic Principles and Concepts II: Immunology 26 Nematodes: Ascaridomorpha, Intestinal Large and Pathology 25 Roundworms 433 4 Parasitic Protozoa: Form, Function, 27 Nematodes: Oxyuridomorpha, and Classification 43 Pinworms 447 5 Kinetoplasta: Trypanosomes and Their 28 Nematodes: Gnathostomatomorpha and Kin 61 Spiruromorpha, a Potpourri 453 6 Other Flagellated Protozoa 89 29 Nematodes: Filaroidea, Filarial Worms 463 7 The Amebas 107 30 Nematodes: Dracunculoidea, Guinea Worms, 8 Phylum Apicomplexa: Gregarines, Coccidia, and Others 479 and Related Organisms 123 31 Phylum Nematomorpha, Hairworms 487 9 Phylum Apicomplexa: Malaria Organisms 32 Phylum Acanthocephala: Thorny-Headed and Piroplasms 147 Worms 495 10 Phylum Ciliophora: Ciliated Protistan 33 Phylum Arthropoda: Form, Function, Parasites 175 and Classification 513 11 Microsporidia and Myxozoa: Parasites with Polar 34 Parasitic Crustaceans 537 Filaments 183 35 Pentastomida: Tongue Worms 561 12 The Mesozoa: Pioneers or Degenerates? 195 36 Parasitic Insects: Phthiraptera, Chewing 13 Introduction to Phylum and Sucking Lice 569 Platyhelminthes 201 37 Parasitic Insects: Hemiptera, Bugs 581 14 Trematoda: Aspidobothrea 211 38 Parasitic Insects: Fleas, Order 15 Trematoda: Form, Function, and Classification Siphonaptera 589 of Digeneans 219 39 Parasitic Insects: Diptera, Flies 601 16 Digeneans: Strigeiformes 247 40 Parasitic Insects: Strepsiptera, Hymenoptera, 17 Digeneans: Echinostomatiformes 265 and Others 627 18 Digeneans: Plagiorchiformes and 41 Parasitic Arachnids: Subclass Acari, Ticks Opisthorchiformes 277 and Mites 639 19 Monogenoidea 295 20 Cestoidea: Form, Function, and Classification of Tapeworms 313 Glossary 661 21 Tapeworms 341 Index 683 22 Phylum Nematoda: Form, Function, and Classification 369 contents Preface xv Basis of Self and Nonself Recognition in Adaptive Responses 30 Antibodies 30 Lymphocytes 31 1 Introduction to Parasitology 1 Subsets of T Cells 31 T-Cell Receptors 32 Relationship of Parasitology to Other Generation of a Humoral Response 32 Sciences 1 Cell-mediated Response 33 Some Basic Definitions 2 Inflammation 34 Interactions of Symbionts 2 Acquired Immune Deficiency Syndrome Hosts 4 (AIDS) 35 Parasitology and Human Welfare 4 Immunodiagnosis 35 Parasites of Domestic and Wild Animals 6 Pathogenesis of Parasitic Infections 36 Parasitology for Fun and Profit 7 Accommodation and Tolerance in the Host- Careers in Parasitology 7 Parasite Relationship 38 References 8 Overview 39 Additional References 9 References
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